Plant Physiology: From Historical Roots to Future Frontiers E-Book
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- Herausgeber: Bentham Science Publishers
- Kategorie: Wissenschaft und neue Technologien
- Serie: Current and Future Developments in Physiology
- Sprache: Englisch
Plant Physiology: From Historical Roots to Future Frontiers provides an in-depth exploration of the principles and advancements in plant physiology. Spanning eleven comprehensive chapters, the book traces the field's historical evolution and covers modern applications such as stress physiology, growth regulators, genomics-proteomics, and bioinformatics. It highlights the integration of cutting-edge technologies like CRISPR-Cas and artificial intelligence, offering insights into their transformative potential in plant science.
Written for a scholarly audience, this book bridges traditional plant physiology with future-oriented innovations, providing a molecular and cellular perspective on growth, metabolism, and physiological processes. It serves as a valuable resource for understanding current challenges and emerging solutions in plant physiology.
Key Features:
- Coverage from historical foundations to advanced research topics.
- Focus on molecular mechanisms and quantitative approaches.
- Discussion of transformative technologies, including CRISPR-Cas and AI.
- Insights into secondary metabolites, stress metabolism, and bioinformatics.
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Seitenzahl: 580
Veröffentlichungsjahr: 2024
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FOREWORD
Current and Future Developments in Physiology (Volume 2) is a book that is published for the benefit of botany. The people spearheading the initial steps of this service hope that this book will serve as a valuable resource for all plant physiologists in publishing important, fundamental discoveries that further our understanding of plant growth, development, and metabolism. The editor and authors of the chapters see their work as providing committed support to the study of plant physiology as a whole.
With the rapid advancement of technology, science has begun to occupy an increasingly larger place in the lives of humanity. Research in plant physiology is increasingly evolving into the fields of artificial intelligence and bioinformatics, which include computerized applications. Concurrently, it constantly investigates in more depth the problems of developmental metabolism under the leadership of physiologists well-trained in molecular approaches, biophysics, and biochemistry methods. In this context, this book combines previous knowledge in the field of basic plant physiology with artificial intelligence and bioinformatics, including biotechnology and molecular biology approaches.
Research is uncovering actual issues that are pressing the greatest intellectual pursuits of humanity to address. Therefore, this book is available for all plant sciences, where physiological approaches must be employed to solve encountered difficulties in order to improve this major field of research overall. In addition to serving as a method for bringing all plant physiologists together into a cohesive, effective working group, my hope is that it will serve as a resource for plant physiologists across all fields of study by offering a central setting in which we can collaborate in the advancement of plant physiology without interfering with other groups' scheduled activities.
I think that this book, which consists of eleven chapters, each containing useful information, will be an even more useful resource with the support and constructive criticism of basic science plant physiologists and applied physiologists from all nations. I would like to express my endless thanks to the editor, book authors, publication editors, graphic designers, and all those who contributed to the preparation and publication of the book. I hope you will read the book as a useful resource.
PREFACE
Plant physiology is a science that studies the symptoms and causes of various vital events that occur throughout the life of plants. The vital events occurring in plants are the result of chemical and physical changes in the living matter of the cell. So, in more general terms, we can define the events that occur as a result of the physical and chemical changes that occur in living things as physiological. Plant physiology tries to answer the question of how and why these physiological events occur in plants, and thus plant physiology reveals the laws and principles in force for physiological events. While presenting these laws and principles, the laws of physics and chemistry are undoubtedly used to a large extent. This reveals that physiology has a close relationship with physics and chemistry. It should also be noted that plant physiology has a special importance in biology because it is a science based on quantitative results, just like chemistry and physical sciences. Because plant physiologists must not only provide descriptive explanations but also explain events with quantitative values.
This book of eleven chapters on classical plant physiology and basic mechanisms aims to present to the reader, by combining many approaches based on the molecular basis of the development of plant physiology, the elucidation of basic metabolic pathways, and the events taking place at the cellular level, starting from the history of plant physiology to current molecular approaches and artificial intelligence-supported applications.
I would like to thank all the book authors who made valuable contributions to the preparation of this book, my advisors Prof. Dr. Fusün GÜMÜŞEL, Prof. Dr. Yelda ÖZDEN ÇİFTÇİ, Dr. Fernanda VİDİGAL DUERTA SOUZA and Dr. Dave ELLIS, who helped me excel in this field and provide useful products, and our families who always supported me.
DEDICATION
To the world you are a mother, but to your children you are the world, Dedicated to my mother and all mothers…
List of Contributors
Historical Development of Plant Physiology
Ergun Kaya1,*Abstract
Although the basis of plant science is identified with the history of humanity, studies in the field of plant physiology based on both the development process of science and technological developments date back to the very recent past. In 1727, English physiologist, inventor, and chemist Stephen Hales published a book called 'Vegetable Statick' and in his book, Hales explained how water is mobilized in plants and laid the foundations of plant physiology. Since then, great developments in technology and biotechnology have allowed plant physiology to grow in a logarithmic manner. Today, many metabolisms have been enlightened both at the cellular level and at the tissue and organ level, and new studies are being added to these studies every day. In addition to the significant advances brought about by technological advancement, research in the fields of nutrition, plant chemistry, particularly in the agricultural sector, and genetics and molecular biology, though often fraught with ethical issues, has produced some truly groundbreaking discoveries. Within this framework, the goal of this chapter is to elucidate the features of the development processes by examining the history of plant biotechnology development, how technological advancements have accelerated this process, and what key studies were conducted during these phases.
INTRODUCTION
Plant physiology investigates the biological events that occur during the life process of a plant, from the beginning as a seed to the stage of producing seeds again. Plant functions are basically based on the principles of physics and chemistry. Plant physiology is studied with the application of modern physics and chemistry techniques to the plants studied, so research in plant physiology also deepens with the advancements in these branches of science [1, 2].
The development of tissues in plants is closely related to the environment in which the plant is located and the physiological events that occur accordingly.
Light, humidity, temperature, water, and gravity are important environmental factors affecting plant growth. Structure and function are closely related; that is, living things come into question because of the coexistence of genes, enzymes, other molecules, organelles, cells, tissues, and organs. For this reason, studies on plant physiology are closely related to plant anatomy, cell biology, and structural and functional chemistry [3, 4].
The target principle of plant physiology is how plants reproduce, grow, and develop. From ancient times to the present, when people started collecting seeds to grow nutritional plants, they saw that plants needed optimum environmental conditions such as sunlight, warmth, and moist soil and that the best-quality seeds produced the most productive plants [5]. They did, however, observe the advantages of practices like fertilization, irrigation, and hoeing. Agricultural activities, which spanned a very long time, enabled the development of new varieties and the cultivation of various species (Fig. 1). The information obtained from basic plant growth and morphological analyses, which are among all these preliminary study activities, laid the foundations of plant physiology [5, 6].
Fig. (1)) Medicinal-aromatic plant species cultivated in the collection garden of Muğla Metropolitan Municipality, Department of Agricultural Services, Garden for Local Seed Center. a.Echinacea purpurea (L.) Moench; b.Salvia fruticosa Mill.Early Experiments on Growth and Development
The first physiological approach to growth was directed at answering the question of where a plant gets the components necessary for its growth. Jean Baptista Van Helmont [7], who lived in the early 1600s, made a suggestion for this approach. According to Helmont, a Belgian doctor, the only source for the growth and development of a plant was water. The researcher irrigated the willow sapling he planted in a pot with only rainwater and grew a huge tree on a very small piece of soil. The researcher knew about CO2 at the time but never anticipated that it could be one of the key growth drivers [7]. Some studies that have provided significant developments in plant physiology are listed chronologically in Table 1, and are briefly summarized. In the 1700s, Antoine Lavoisier discovered that the matter resulting from organic synthesis was composed of oxygen and carbon on a large scale [8].
Joseph Priestley, Jan Ingenhousz, and Jean Senebier showed that plant leaves take up carbon dioxide in light and emit an equivalent amount of oxygen [9, 10]. Later, Nicholas de Saussure noted the involvement of water in the process. In the dark, the opposite happened: plants breathed like animals, taking in oxygen and releasing carbon dioxide [11, 12]. Julius Robert von Mayer observed that the process converts light energy into the chemical energy of organic carbon. Thus, the growth of seedlings in the dark or roots in the soil came at the expense of this energy. Therefore, in the 19th century, photosynthesis, although not understood biochemically, was considered the primary and fundamental synthetic process in plant growth [13].
Previous Studies in Plant Nutrition and Transport
In the 1830s, Justus von Liebig proposed the Law of Minimums. In this law, it is emphasized that the growth and development of plants are limited by the mini- mum amount of nutrients in the soil, and this view has been confirmed by experiments conducted around the world [14, 15].
One of the first studies on the uptake of nutrients and mineral substances from the soil in plants was carried out by Julius Sachs and his colleagues in order to determine that nitrogen, potassium, phosphate, sulfur and other elements in quantitative terms are of great importance in small soil components for plant growth, and they used various chemical analyses in these studies [14]. Due to the presence of inorganic nutrients, particularly nitrogen, in the soil, fertilization has long been recognized as being important. They can be introduced to the soil as inorganic salts, like potassium nitrate, . The organic matter of manure, or the residue of its decay, has been shown to contribute to improving the soil or soil structure. From these discoveries, different approaches to the modern agricultural use of chemical fertilizers have been introduced [14, 16].
One of the pioneering studies on water uptake from the soil in plants was carried out by Van Helmont on the willow plant. In this study, metabolic processes in the plant were observed by continuously watering it with much more water than the tree contains [17]. In 1727, Stephen Hales, an English clergyman, and amateur physiologist, published his work titled “Vegetative Statics” as a result of his studies on transpiration [18], plant growth, and gas exchange in plants. Hales suggested that water coming from the soil moves towards the leaves, where it is lost as water vapor through a process called transpiration. Subsequent research in the nineteenth and early twentieth centuries showed that water diffuses through stomata, and pores in the leaf epidermis [17, 18].
The opening and closing mechanism of stomata is based on the principle of stomata taking in and giving out water, depending on the presence or absence of water. Capillary forces occurring in the vascular bundles along the mesophyll layer in leaves, with some contribution from osmosis, pull water columns up through the open vessels and tracheids of the xylem that carry nutrients from the roots. The harmony between water molecules and their adhesion to cell walls prevents the tense water columns from breaking even in very tall trees (adhesion-cohesion force of water). This mechanism was first proposed by Henry Dixen and John Joly in 1895, and in the twentieth century, numerous researchers confirmed and developed this “adaptation-tension” theory of transportation [19, 20].
Stephen Hales also measured the root pressure of plants whose apical meristematic region had been cut. Subsequent studies have shown that under conditions of quality soil, appropriate moisture, and aeration, roots actively secrete high concentrations of salt into the root xylem, creating a high osmotic pressure (guttation) that forces water up the stem and out through pores at the tips of the leaves. In 1926, E. Munch proposed a similar mechanism for translocation, which is the transfer of sugars from leaves to roots and other plant parts via the phloem. This mechanism is now known as the pressure flow model [21, 22].
Cellular and Molecular Plant Physiology
As the twentieth century approached, plant physiologists increasingly turned to chemistry and physics to answer fundamental questions. In addition, they created their groups and published new journals with the study's findings. This has had a significant impact on increasing the level and amount of research. It has been found that many of the results obtained from more comprehensive medical, animal, and microbiological research on the basic biochemistry of cell growth and function also apply to plant cells. Anatomical studies have yielded structural details to support physiological findings, and microscopic cell structure has been revealed by electron microscopy [23, 24].
Environmental, hormonal, cellular and genetic factors on growth and development have been studied in detail, but there are many mechanisms yet to be learned. For instance, plants create ethylene, a basic two-carbon gas that controls seed germination and begins the ripening of fruit. It has been discovered that auxin or cell growth hormone translocation causes phototropisms and geotropisms [25]. In some cases, auxins can also stimulate cell division. Other plant growth regulators, gibberellins, regulate cell division in the root tip and activate enzyme formation in seed germination [25, 26].
The development of plant tissue culture technology in the early 1900s led to the discovery of cytokinins, which are growth regulators effective in the division of plant cells [27]. In 1902, Gottlieb Haberlandt cultured the plant cells he isolated in a simple nutrient solution and presented this initiative as a classic article at the meeting of the Vienna Academy of Sciences in Germany [28]. In 1935, Snow demonstrated the role of indole acetic acid (IAA) in promoting cambial activity for rapid cell proliferation [29]. In 1943, Philip R. White introduced a synthetic plant tissue culture medium composition (White medium-WM) containing vitamin B, and tomato root cultures could be preserved in this environment for as long as 30 years [28]. In 1962, Toshio Murashige and Folke K. Skoog reported the plant tissue culture medium composition (Murashige-Skoog medium-MS medium) that has been widely used until today. This initiative is important in that it gives insight into the potential of the cell and demonstrates a new technology as plant tissue culture. Abscisic acid, another plant growth regulator, initiates the senescence and shedding of leaves in autumn and causes stomata to close under water stress. Today, studies on growth regulatory compounds are continuing, but there are still points that need to be clarified on the basic mechanisms of the interaction of these molecules with each other [28, 30].
Photoperiodism, the regulation of flowering according to day length, was discovered by Wightman W. Garner and Harry A. Allard in 1920. Further studies were found in the 1930s by German biologist Erwin Bünning that sleep movements, such as the drooping of bean leaves in the evening, were controlled not by the onset of darkness but by a biological “clock”, a circadian rhythm. Phytochrome was discovered by Harry Borthwick in 1952 and was found to be the pigment central to photoperiodism [31, 32].
In recent years, molecular biology studies have gained great momentum in attempts to find genes responsible for physiological processes and metabolic pathways. The structure of chlorophyll in photosynthesis has been determined and it has been revealed that it is localized in the inner membranes of the chloroplasts of mesophyll cells. The finding that the red and blue portions of the light spectrum are effective led to the discovery that two light reactions are required [33]. In 1929, Cornelis Bernardus van Niel used radioactive water to show that water, not carbon dioxide, was the source of the oxygen released during photosynthesis [34, 35]. Sugar was found to be synthesized in the stroma of the chloroplast, and the molecular details of its formation were solved by Melvin Calvin and Andrew Benson in 1963. All plant cells have been found to respire. This is understood to be an energy-yielding process that involves a different membranous organelle, the mitochondria, which is essentially the same as in animals and yields metabolic energy available for transport reactions and synthesis of cell material [36].
The formation of fats and oils from carbohydrates was found to be similar to that in animals, but plants had the additional ability to convert fats in germinating seeds into carbohydrates such as glucose, which are used in cellulose wall formation. The symbiotic relationships of plants and microorganisms were investigated, especially in cases of reduced nitrogen formation from atmospheric nitrogen by nodule bacteria. In this way, various types of energy metabolism that occur in prokaryotic organisms were discovered, and inferences could thus be made about the role that specific microorganisms play in the cycling of elements. In addition, it became possible to elucidate many details about metabolic pathways and their regulation [37, 38].
Described by Friedrich Laibach in 1943 as a model organism for developmental and physiological analyses, Arabidopsis thaliana became central to scientists' efforts to understand plant genomes at the end of the twentieth century. The complete sequence of this genome was elucidated in 2000 by an international consortium of plant geneticists. Following the introduction of recombinant DNA technology into the scientific world, the acceleration of technological developments and the spread of DNA/Protein-based techniques after the twentieth century allowed studies to elucidate many cellular metabolic pathways to continue rapidly [39, 40].
Programmed Cell Death or Apoptosis
The process of planned cell death is called apoptosis. It is applied in the early stages of development to get rid of undesirable cells, like the ones that grow between a developing hand's fingers. Adults employ apoptosis to get rid of cells that are too damaged to be repaired [41]. In plants, as in animals, programmed cell death or apoptosis is the final stage of genetically controlled cell differentiation. In some cases, they acquire special functions as they die (such as vascular tissues, and fibers) or, conversely, cells die after completing their duties. This type of programmed cell death is called developmental cell death and is controlled by an endogenous program. The other type of cell death occurs as a result of different environmental signals or pathogen attacks, including biotic and abiotic stimuli, and they cause changes in the original cell program. Senescence, also referred to as biological aging, is the progressive loss of functional traits in living things. In the latter stages of an organism's life cycle, at least, whole organism senescence refers to a rise in mortality or a fall in fecundity with advancing age. Senescence is often associated with apoptosis. Programmed cell death in plants occurs in response to specialized conditions involving the hypersensitivity response to pathogens and the development of tracheal elements. It may differ from programmed cell death because senescence can be delayed and reversible. This indicates that programmed cell death is a non-essential part of the senescence process [41, 42].
The Age of Artificial Intelligence in Plant Physiology
Today, artificial intelligence technologies (Fig. 2). operate in many areas, from automatic editing applications on mobile phones to facial recognition, using satellite systems and precision agriculture applications. Although the use of these technologies does not have a long history, they have recently enabled serious developments in many fields. Increasing computing power, increasing ability to collect, manage and store huge amounts of data, and algorithmic advances are the most obvious examples. Multiple applications of artificial intelligence have been developed, each with their own techniques, strengths, and weaknesses, making certain approaches a better match for certain problems than others [43, 44].
Fig. (2)) Schematic representation of the relationship between artificial intelligence, machine learning, and data science [45].The use of artificial intelligence neural networks (e.g., deep learning) forms the basis for rapid and effective machine learning in solving a number of scientific problems in plant science and many physiological problems. For example, deep learning technologies have recently been useful in a variety of predictive tasks, such as species identification, determination of natural ranges, modeling of plant species distributions, diagnosis of diseases, and identification of disease agents. In addition, studies on comparative genomics and gene expression have been highly effective in understanding metabolic pathways and physiological activities in the cell. Moreover, new approaches to combining high-resolution imaging technologies with artificial intelligence and machine learning and digital repositories are now poised to revolutionize the study of plant phenology and functional traits. Artificial intelligence technologies are advancing rapidly due to their advantages in plant physiology and cellular metabolic processes, combining the development of alternative approaches best suited to specific questions, data sources and analytical techniques [46, 47].
CONCLUSION
Biological sciences are developing very rapidly, with the help of technological developments such as genomics, molecular genetics, high-throughput genomes, and image analysis in the post-genomic era we are in. As a result, unknowns regarding the structure, function, and development of plants are rapidly being clarified. Especially since the early 2000s, our knowledge about the relationships between signal pathways and gene networks that complement each other and work in these characteristics of plants has been increasing day by day. Considering that the metabolism, growth, and development of the plant are a whole and continuous interaction, many unexplained mechanisms, concepts, and metabolic pathways in plant physiology will be elucidated in the future, thanks to the methods updated with the developing technology.
The ever-increasing world population confronts agriculture with new challenges, which necessitates greatly increasing the productivity of future agriculture. To continue to increase crop yields sufficiently in the future, a new green revolution seems inevitable. However, as the improved yields achieved gradually increase, conventional breeding is increasingly reaching its limits but is unlikely to keep pace with the rate of population growth in the long term.
Advanced technologies and molecular plant physiology approaches have now begun to open new horizons in the fight against an ineffective process in the plant that competes with photosynthesis. This refers to the plant's respiratory processes (photorespiration) that cause energy loss and therefore limit growth, and researchers want to modify metabolic pathways or develop new ways to bypass or improve photorespiration. This suggests that this would be possible, for example, by changing enzyme properties or adding new proteins.
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Ergun Kaya1,*,Selin Galatalı1Abstract
Plants often encounter environmental stressors in both wild and cultivated environments. Certain environmental stressors, like air temperature, only last a few minutes, but others, like soil water content, might persist for several days. Stress might last for months if there is a mineral shortage in the soil. This chapter gives an overview of the ways that soil, climate, and stress affect the spread of different plant species. Thus, it is crucial for agriculture and the environment to comprehend the physiological mechanisms that underlie plants' methods of adaptation and acclimatization to environmental challenges. A common definition of stress is an outside influence that negatively impacts plants. Stress tolerance and the concept of stress are closely related. The capacity of a plant to withstand adverse environmental conditions is known as stress tolerance. One plant may not find stress in the same environment as another. Based on the fundamental ideas of stress physiology in plants, this chapter seeks to provide a modern and fundamental explanation of the metabolic processes that occur in cells.
INTRODUCTION
Living things, by their nature, are in constant contact with the external environment. If inappropriate conditions occur in their environment, they are exposed to stress conditions due to lack of adaptation. When environmental conditions change so much that they negatively affect the normal growth and development of a plant, the situation that occurs in the plant is called stress. In other words, it is defined as external factors that have negative effects on the plant. In many cases, stress is a concept that needs to be explained by relating it to the survival of the plant, its ability to produce products, biomass accumulation and assimilation [1, 2].
As a matter of fact, abiotic stresses such as drought, salinity, extreme temperatures, chemical toxicity, and oxidative stress are serious threats that disrupt agricultural activities and deteriorate the environment. For example, abiotic stress is the primary cause of crop yield loss worldwide, greatly reducing average crop yields in the most productive crop plants. Accumulated data on how plants respond to biotic and abiotic stress and how stress affects the developmental processes in the plant life cycle have enabled the development of new approaches on the subject [3, 4].
Plants encounter many stress factors during their lives. According to Levitt, stress factors are divided into two: biotic and physicochemical [5]. Biotic factors include stress factors caused by the infection of microorganisms (fungi, bacteria, and viruses) and attacks by harmful animals. Abiotic factors are environmental factors such as water, temperature, radiation, chemicals, magnetic and electrical fields [6]. Plants, which do not have the option of avoiding the stressor by moving away from it due to their sessile nature, are directly exposed to stress, unlike animals. This direct effect negatively affects growth and development and causes plant organs to lose their vitality [7, 8].
Damage caused by stress factors varies depending on the plant type, tolerance, and adaptation ability [9, 10]. Considering that plants encounter many stress factors in nature throughout their lives, it is very important to elucidate stress-related mechanisms and develop tolerant species and varieties. For this purpose, in this chapter, the molecular and biochemical events that occur in plants under stress conditions will be discussed and the responses to stress will be tried to be explained [7, 11].
RESISTANCE TO WATER SHORTAGE AND DROUGHT
Drought resistance mechanisms are divided into several types. First, postponement of drying (the ability to retain water in the tissue) and tolerance to drying. These two are sometimes referred to as drought tolerance at high and low water potentials, respectively. A third mechanism is escape from drought. This mechanism involves completing the life cycle during the rainy season, before drought occurs. There are two categories of people who delay drying: those who do not waste water and those who do. Individuals who do not waste water use it in moderation. In order to use it later in life, these plants store part of the water in the soil. Individuals that wastewater consume a lot of water. The mesquite tree (Prosopis glandulosa Torr.) is an example of a water waster. This plant, whose roots can reach very deep, has destroyed the semi-arid grasslands in the southwestern United States [12]. Since it uses excessive water, it prevents grasses of agricultural value from settling in that area. A plant with a high ability to gain and use water will be more resistant to drought [12, 13].
Some plants have adaptations such as the C4 and CAM photosynthesis pathways (Fig. 1). These adaptations allow plants to use more water. Additionally, plants have acclimation mechanisms that are activated to respond to water stress. Water scarcity is defined as any water content of a tissue or cell below the highest water content in the plant. Water stress has some effects on growth: it is the limitation of leaf expansion (i), water shortage stimulates leaf abscission (ii), during water scarcity, roots grow towards moist regions deep in the soil (iii), stomata close in response to abscisic acid during water scarcity (iv), water shortage limits photosynthesis (v), osmotic stress promotes Crassulacean Acid Metabolism (CAM) in some plants (vi), and leads to changes in gene expression (vii) [14, 15].
Fig. (1)) Types of photosynthesis in C4 and CAM [18].Water Scarcity Limits the Leaf Expansion
As the water content of the plant decreases, the cells shrink and the cell walls become looser. This decrease in cell volume causes the turgor pressure in the cells and subsequently the solute concentration to decrease. As the area it covers decreases, the plasma membrane thickens and the pressure on it increases. Decreased turgor is the first and most important biophysical effect of water stress. Inhibition of cell expansion in the early stages of water scarcity slows leaf expansion. As the leaf area decreases, water loss through transpiration decreases. Thus, the limited amount of water in the soil is effectively preserved for a long time. Therefore, the reduction in leaf area can be considered the first line of defense against drought [16, 17].
Water Shortage Stimulates Leaf Abscission
Water has a significant impact on the physiological and morphological development of plants. Water stress can also affect plants. Plants may organalise in response to water stress in order to preserve the dynamic water balance. The total leaf area of a plant (number of leaves × surface area of each leaf) does not remain constant after all the leaves have matured. If plants are exposed to stress after the leaf area has been formed, the leaves will turn yellow and eventually fall off [19, 20].
Root Growth Response to Moisture During Water Scarcity
One significant environmental element limiting crop yields and plant development is limited water availability. The ability of plants to maintain root growth in order to have continuous access to soil water is one of their most notable responses to water shortages. The innate complexity of root systems and their interactions with the soil environment has hindered understanding advances even though the adaptive significance of maintaining root growth under water deficiencies was recognized early on. Moderate water scarcity also affects the development of the root system. The ratio of the mass of the root to that of the trunk is determined by the functional balance between water uptake from the roots and photosynthesis by the above-ground parts [21, 22].
Abscisic Acid-Induced Stomatal Closure During Water Scarcity
Closing of stomata constitutes another line of defense against drought. The uptake and loss of water into the guard cells changes the turgor of these cells, allowing the stomata to open and close. Since guard cells are located in the epidermis, they lose their turgor by losing water directly to the atmosphere through evaporation. The decrease in turgor causes the stomata to close hydropassively. A second mechanism, called hydroactive closure, allows the stomata to close when all leaves and roots lose water. Hydroactive closure depends on metabolic processes in guard cells. The decrease in the amount of dissolved substances in the guard cells causes water loss and turgor decrease and causes the stomata to close. Therefore, the hydraulic mechanism of hydroactive closure is the opposite of the stomatal opening mechanism. However, hydroactive closure is controlled differently than stoma opening. Decreasing the water content of the leaf initiates the loss of dissolved substances from the guard cells. Abscisic acid (ABA) plays a major role in this process. ABA (Fig. 2)., which is synthesized at a low rate and continuously in mesophyll cells, tends to accumulate in chloroplasts. When mesophyll cells lose moderate amounts of water, two events occur: (a) Some of the ABA accumulated in chloroplasts is released into the apoplast (cell wall space) of the mesophyll cell. The unequal distribution of ABA depends on pH gradients within the leaf, the weakly acidic nature of the ABA molecule, and the permeability of cell membranes. The unequal distribution of ABA ensures that a portion of ABA is transported to guard cells by transpiration flow. (b) ABA is synthesized at a higher rate and more ABA accumulates in the leaf apoplast. The increase in ABA concentration as a result of rapid synthesis increases or prolongs the previously formed sealing effect of accumulated ABA [23, 24].
Fig. (2)) Function of ABA in stomatal defense against water shortage [25].Water Shortage Limits Photosynthesis
Photosynthesis is much less sensitive to turgor change than leaf expansion. Therefore, the rate of photosynthesis in the leaf (expressed per unit leaf area) does not respond as well to leaf expansion to moderate water stress. However, moderate water stress generally affects both photosynthesis and stomatal conductance. In the early stages of water stress, water use efficiency may increase as stomata close (i.e., more CO2 can be taken in per unit amount of water lost by transpiration); because stomatal closure inhibits transpiration more than intercellular CO2 concentration [26, 27].
Osmotic Stress Promotes Crassulacean Acid Metabolism
CAM can be induced by salt or water stress, which turns on gene expression for the manufacture of the component enzymes. CAM is also a plant adaptation in which stomata are kept open at night and closed during the day. The difference between the vapor pressure of the leaf and the air, which enables transpiration to occur, decreases significantly when both the leaf and the air cool. Therefore, the water useability of CAM plants is the highest of all measured. CAM is very common in succulent plants such as cacti. Some succulent species show facultative CAM properties. That is, they turn to GLASS when exposed to water shortage or saline conditions [28, 29].
Osmotic Stress Alters Gene Expression
As previously noted, the accumulation of compatible solutes to respond to osmotic stress requires the activation of metabolic pathways that enable the synthesis of such substances. Some genes encoding enzymes responsible for osmotic regulation are switched on under the influence of osmotic stress and/or salinity and cold stress. These genes encode the following enzymes: Δ'1-Pyrroline-5-carboxylate synthase (i), a key enzyme in the proline biosynthesis pathway; Betaine aldehyde dehydrogenase (ii), which is involved in the accumulation of glycine betaine; Myo-Inositol 6-O-methyltransferase (iii), a rate-limiting enzyme in the accumulation of pinitol; Glyceraldehyde-3-phosphate dehydrogenase (iv), which is expressed more during osmotic stress [30, 31].
Other genes regulated by osmotic stress encode proteins involved in membrane transport. ATPases and channel proteins that allow water to pass through, aquaporins, are among these. Additionally, stress stimulates some protease enzymes. These enzymes can break down (remove and recycle) other proteins whose structure is disrupted during stress. The protein called ubiquitin labels target proteins for proteolytic degradation. In Arabidopsis [31], mRNA synthesis responsible for ubiquitin increases after drought stress. Additionally, some osmotically stimulated heat shock proteins protect or renaturate proteins inactivated by drying [32, 33].
HEAT STRESS AND HEAT SHOCK
Most of the tissues of higher plants cannot survive prolonged temperatures above 45°C. Growth-arrested cells and dehydrated tissues (e.g., seeds and pollen) can survive at much higher temperatures than growing vegetative cells that contain water. While actively growing tissues do not survive temperatures above 45°C, dry seeds can withstand 120°C and pollen of some species can withstand 70°C. In general, only single-celled eukaryotes can complete their life cycles at temperatures above 50°C, and only prokaryotes can divide and grow at temperatures above 60°C. Periodic application of short-term sublethal heat stress often stimulates tolerance to lethal temperatures. We refer to this phenomena as induced thermotolerance [34, 35].
High Leaf Temperature and Water Shortage Lead to Heat Stress
Many CAM and succulent higher plants, such as Opuntia and Sempervivum, are adapted to high temperatures. These plants can withstand tissue temperatures of 60-65°C under intense sunlight in summer. Since CAM plants close their stomata during the day, they cannot cool down through transpiration. Instead, they radiate heat from sunlight by reflecting back long wavelength (infrared) rays and also lose heat by conduction and convection [36, 37].
The increase in leaf temperature during the day is striking in plants of arid and semi-arid climate regions where drought and sunlight are intense. Heat stress is also a potential hazard in greenhouses. Because air movement in greenhouses is slow and humidity is high, leaf cooling decreases. Moderate heat stress slows the growth of the entire plant [38, 39].
At High Temperatures, Photosynthesis is Inhibited Before Respiration
The unique change in the redox state linked to thylakoid membranes is indicative of a cellular energy imbalance induced by environmental stress, which is sensed globally by photosynthesis. Because photosynthesis is extremely vulnerable to stress from high temperatures, it frequently stops before other cell activities are compromised. The ability of maximum plants to precisely adjust their photosynthetic traits to their growing temperatures is demonstrated in great detail. The most notable observable fact is that plants can increase their photosynthetic efficiency at their new growth temperature by shifting the optimal temperature of photosynthesis in response to changes in growth temperature or seasonal temperature swings. Both photosynthesis and respiration are inhibited at high temperatures; however, as temperature increases, photosynthesis rates decrease before respiration rates. The temperature at which the amount of CO2 fixed by photosynthesis is equal to the amount of CO2
